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Copyright © 2021 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4927.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:9A141A50-151D-4261-9A62-FD300B521E86
Abstract
Global trade has increased the invasion risk of exotic organisms and damaged agricultural and natural ecosystems. The
Bureau of Animal and Plant Health Inspection and Quarantine (BAPHIQ) handles quarantine services of animal- and
plant-associated pests and diseases in Taiwan. The predatory mite family Phytoseiidae (Acari: Mesostigmata) is a well-
known group due to the potential use of certain species as biocontrol agents for small phytophagous pests. Some species
are available in commercial markets and frequently used in biological control in many agricultural systems, especially
in greenhouse crops. However, exotic biological control agents may interfere with natural or naturalised populations of
predatory mites and they may threaten indigenous populations via intraguild predation. The present study aims to provide
a checklist of phytoseiid mite species found in plant quarantine from 2006–2013. Twenty-five species belonging to two
subfamilies and eight genera were found in samples imported to Taiwan from twelve countries, including one new species
Typhlodromus (Anthoseius) ueckermanni sp. nov. from South Africa. The checklist provides distribution, remarks, and
also an identification key for all species.
Introduction
International trade has allowed exotic species to move across geographical barriers and invade to new areas. Exotic
organisms may have negative effects on agricultural and natural ecosystems (Hulme 2009). Numerous exotic natu-
ral enemies have been introduced and provided successful pest control. However, few studies have focused on the
negative effects of introduced biological control agents (van Lenteren et al. 2003). The family Phytoseiidae (Acari:
Mesostigmata) is a famous group in terms of their potential as biological control agents, and contains more than
2,700 species (including synonyms) worldwide (Chant & McMurtry 2007; Demite et al. 2020). Certain phytoseiids
are available in commercial markets to be used in biological control of spider mites, eriophyid mites, and small soft
bodied insects such as thrips and whiteflies, and thus provide a valuable contribution to the agroecosystem (Mc-
Murtry et al. 2013). However, exotic biological control agents may interfere with natural or naturalised populations
of predatory mites and they may threaten indigenous populations via intraguild predation. For example, Harmonia
axyridis (Pallas) is a famous generalist predator for hemipteran pests, however, now it is considered as an invasive
species in many countries where it has been spreading rapidly and declined the populations of native species since
the early years of the 21st century (Brown et al. 2018). In addition, Sato & Mochizuki (2011) reported that two
exotic phytoseiids (Neoseiulus cucumeris and Amblyseius swirskii) were introduced to Japan, have advantage over
an indigenous species, Gynaeseius liturivorus (Ehara) via intrguild predation. In this regard, the identification of
predatory mite species in plant quarantine samples imported from foreign countries is crucial for the evaluation of
their environmental risk over natural population.
This study provides a checklist and an identification key of 25 phytoseiid species found during plant quarantine
Mite specimens examined in this study were collected in plant quarantine by inspectors of Bureau of Animal and
Plant Health Inspection and Quarantine during 2006–2013. Specimens were mounted in Hoyer’s medium, examined
under an optical microscope (Olympus BX51). Measurements were taken using a stage-calibrated ocular microm-
eters and ImageJ 1.47 (Schneider et al. 2012). All measurements were provided in micrometres (μm) and holotype
measurements are shown in bold type for new species, followed by their mean and range (in parenthesis). The dorsal
shield lengths were measured from anterior to posterior margins along the midline and the widths measured at j6
and S4 levels. The sternal shield lengths and widths were taken from anterior to posterior margins along the midline
and at st2 level, respectively. The genital shield widths were taken from st5 level. The ventrianal shield lengths were
taken from anterior to posterior margins along the midline including cribrum and the shield widths measured at ZV2
and anus levels. The general terminology used for morphological descriptions in this study follows that of Chant &
McMurtry (2007). The notation for idiosomal setae follows that of Lindquist & Evans (1965) and Lindquist (1994),
as adapted by Rowell et al. (1978) and Chant & Yoshida-Shaul (1991, 1992). The notation for solenostomes and lyr-
ifissures is based on Athias-Henriot (1975). Type specimens were deposited in NTU (Nataional Taiwan University,
Taipei City, Taiwan), remaining specimens in TARL (Taiwan Acari Research Laboratory, Taichung City, Taiwan).
Results
Identification key to phytoseiid mites in plant quarantine in Taiwan based on female specimens
Material examined. Netherlands: one female (HAL095F301) from Chrysanthemum morifolium (Asteraceae), 27
Aug 2006; one female (HAL095F062) from Symphoricarpos albus (Caprifoliaceae), 29 Oct 2006; one female
(HAL100B070) from Skimmia sp. (Rutaceae), 20 July 2011; two females (HAL101B145) from Rosa hybrida (Ro-
saceae), 20 Feb 2012; two females (HAL098B508) from Symphoricarpos sp. (Caprifoliaceae), 20 Sept 2009; one
female (QAR102H025) from Bouvardia sp. (Rubiaceae), 22 June 2013; one female (QAR102H043) from Sym-
phoricarpos sp. (Caprifoliaceae), 8 Sept 2013; three females (QAR102H045) from Bouvardia sp. (Rubiaceae), 8
Sept 2013.
USA: two females (HAL099C254) from Malus pumila (Rosaceae), 25 Nov 2010.
Canada: one female (QAR101H027) from Capsicum annuum (Solanaceae), 28 Sept 2011.
Japan: one female (QAR101H019) from Capsicum annuum (Solanaceae), 12 Sept 2011.
Previous records. Africa: Algeria (Athias-Henriot 1958), Morocco (Tixier et al. 2003).Asia: Azerbaijan (Ab-
basova 1972), Cyprus (Papadoulis et al. 2009), Japan (Chant 1959), Syria (Barbar 2013), Turkey (Çobanoðlu 1991).
Europe: Austria (El-Borolossy 1989), Bosnia and Herzegovina (Döker et al. 2019), Czech Republic (Kabíček
2004), Denmark (Hansen & Johnsen 1986), England (Chant & Hansell 1971), France (Athias-Henriot 1962), Geor-
gia (Wainstein & Vartapetov 1973), Germany (Westerboer & Bernhard 1963), Greece (Papadoulis & Emmanouel
1991), Hungary (Bozai 1980), Italy (Castagnoli et al. 1984), Lativia (Salmane 2003), Moldova (Wainstein 1973), the
Netherlands (Yoshida-Shaul & Chant 1995), Poland (Boczek 1964), Portugal (Carmona 1966), Serbia (Radivojević
& Petanović 1984), Slovakia (Jedlickova 1993), Slovenia (Bohinc et al. 2018), Spain (Villaronga & Garcia Mari
1988), Sweden (Steeghs et al. 1993), Switzerland (Baillod & Venturi 1980), Ukraine (Kolodochka 1973). North
America: Canada (Chant 1957), USA (Chant 1959).
Remarks. The species is distributed worldwide, except in Oceania and South America, and has no previous
record of existing in Taiwan. EPPO (2020) listed this species as a commercially used biological control agent. Most
of specimens came from the Netherlands. However, one specimen originated from Japan. Toyoshima et al. (2016)
recorded this species in Japan for the first time. Further confirmation is required to determine whether this species
was introduced as a commercial product or it is a native population.
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 303
Amblyseius largoensis (Muma, 1955)
Material examined. USA: one female (HAL100B062) from Persea americana (Lauraceae), 19 May 2011.
Previous records. Africa: Angola (Carmona 1968), Benin (Zannou et al. 2007), Ivory Coast (Moraes et al.
1989a), Kenya (Swirski & Ragusa 1978), Mozambique (Rodrigues 1968), Sierra Leone (Zannou et al. 2007), Tanza-
nia (El-Banhawy & Abou-Awad 1990). Asia: China (Chen et al. 1980), India (Gupta 1978), Iran (Daneshvar 1980),
Malaysia (Ehara 2002a), Oman (Hountondji et al. 2010), the Philippines (Corpuz & Rimando 1966), Saudi Arabia
(Alatawi et al. 2017), Singapore (Corpuz-Raros 1995), Sri Lanka (Moraes et al. 2004b), Taiwan (Ehara 1970),
Thailand (Ehara & Bhandhufalck 1977), Turkey (Çobanoðlu 1989b), Vietnam (Fang et al. 2020). Central America:
Cuba (Rodriguez et al. 1981), Dominican Republic (Ferragut et al. 2011), Guatemala (Chant 1959), Mexico (Chant
1959), Jamaica (Denmark & Muma 1978), Puerto Rico (De Leon 1965). Europe: Georgia (Wainstein & Vartapetov
1973). North America: USA (Muma 1955). Oceania: Australia (Collyer 1980), Fiji (Collyer 1980), Hawaii (Prasad
1968), New Caledonia (Schicha 1981b), New Zealand (Collyer 1964b), Papua New Guinea (Schicha & Gutierrez
1985), Vanuatu (Schicha 1981b). South America: Brazil (Ehara 1966), Colombia (Moraes & Mesa 1988), Guyana
(De Leon 1966), Trinidad (De Leon 1967), Venezuela (Aponte & McMurtry 1993).
Remarks. The species is distributed worldwide; however, it is doubtful if all recorded specimens are real A.
largoensis (see Döker et al. 2020). Many researchers (e.g., Liao et al. 2020) indicated that the species can be iden-
tified by the parallel tubular calyx of the spermatheca. Additionally, this species is dominant in southern Taiwan,
especially in the tropical region. This species has also been studied in terms of its potential in biological control of
red palm mite, Raoiella indica Hirst (e.g., Mendes et al. 2018).
Material examined. USA: one male (HAL095F293) from Prunus persica (Rosaceae), 6 July 2006; one female
(HAL095F299) from P. persica (Rosaceae), 24 Aug 2006; two females one male (HAL099C146) from P. persica
(Rosaceae), 10 Sept 2010; one female (QAR101H005) from P. persica (Rosaceae), 22 June 2011; one female (QA-
R101H003) from P. persica (Rosaceae), 4 June 2011; one female two males (QAR101H004) from Fragaria anan-
assa (Rosaceae), 7 June 2011; one female (QAR101H009) from P. persica (Rosaceae), 19 July 2011; one female
(QAR101H010) from P. persica (Rosaceae), 21 July 2011; one male (QAR101H007) from P. persica (Rosaceae), 7
July 2011; one female (HAL101B060) from P. persica (Rosaceae), 29 Sept 2011; one female (HAL101B192) from
P. persica (Rosaceae), 29 June 2012; four females (QAR102H020) from Asparagus officinalis (Asparagaceae), 11
June 2013; one female (QAR102H024) from P. persica (Rosaceae), 21 June 2013; three males (QAR102H026)
from P. persica (Rosaceae), 1 July 2013; one male (QAR102H027) from P. persica (Rosaceae), 17 July 2013; one
female (QAR102H028) from P. persica (Rosaceae), 17 July 2013.
Previous records. North America: Mexico (De Leon 1961), USA (present study).
Remarks. This species is distributed only in Mexico; however, 17 samples examined in this study were col-
lected from the United States; the distribution of the species requires further confirmation. Additionally, most of the
specimens were obtained from Prunus persica.
Material examined. Thailand: one female one male (HAL098B608) from Piper nigrum (Piperaceae), 30 Nov
2009; one female (HAL099B097) from Aranthera beatrice (Orchidaceae), 6 Jan 2010; one male (HAL100B201)
Material examined. Japan: one female (QAR101H028) from Perilla frutescens (Lamiaceae), 28 Sept 2011.
Previous records. Asia: China (Zhu & Chen 1980), Japan (Ehara 1959), South Korea (Ryu & Kim 1998).
Remarks. This species is distributed in East Asia, especially Japan, and is a major predator of spider mites in
orchards in Japan. It is considered as Type III generalist predators (Funayama & Sonoda 2014). The food sources of
the species ranges from spider mites, thrips, whiteflies, pyralid moths, and pollens (Yang et al. 2019).
Material examined. Thailand: one female (HAL095F297) from Asparagus officinalis (Asparagaceae), 15 Aug
2006; one female (HAL095F298) from Ocimum sanctum (Lamiaceae), 21 Aug 2006; one female (HAL101B144)
from A. officinalis (Asparagaceae), 13 Feb 2012; one protonymph (QAR101H015) from A. officinalis (Asparaga-
ceae), 30 Aug 2011; Thailand, one female (QAR101H022) from A. officinalis (Asparagaceae), 17 June 2013.
Netherlands: one female (HAL095F309) from Lactuca sativa var. capitata (Asteraceae), 14 Sept 2006; Neth-
erlands, one female (QAR101H038) from Thymes sp. (Lamiaceae), 5 Sept 2013.
Israel: one female (HAL095G063) from Kochia sp. (Amaranthaceae), 31 Oct 2006; three females (HAL095G064)
from Limonium sp. (Plumbaginaceae), 31 Oct 2006.
Japan: one female (HAL100B075) from Vitis vinifera (Vitaceae), 2 May 2011; one female (QAR101H013)
from Perilla frutescens (Lamiaceae), 23 Aug 2011; one female (QAR101H023) from Chrysanthemum sp. (As-
teraceae), 21 Sept 2011; one female (QAR101H050) from P. frutescens (Lamiaceae), 13 Sept 2013; one female
(QAR101H036) from P. frutescens (Lamiaceae), 3 Sept 2013.
New Zealand: one female (QAR101H001) from Malus pumila (Rosaceae), 5 Apr 2011.
USA: one female (QAR101H014) from Anthriscus cerefolium (Apiaceae), 27 Aug 2011
Peru: six females (QAR101H029) from A. officinalis (Asparagaceae), 21 July 2013;
Previous records. Africa: Algeria (Athias-Henriot 1961), Benin (Zannou et al. 2006), Burundi (Zannou et
al. 2006), Canary Islands (Ferragut & Peña-Estévez 2003), Cape Verde (Ueckermann 1992), Egypt (Abo-Shnaf &
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 305
Moraes 2014), Ghana (Moraes et al. 1989a), Guinea (Ragusa & Athias-Henriot 1983), Kenya (Zannou et al. 2006),
Malawi (Zannou et al. 2005), Morocco (Kreiter et al. 2004), Mozambique (Zannou et al. 2006), Nigeria (Moraes
et al. 1989a), Senegal (Kade et al. 2011), South Africa (Ueckermann & Meyer 1988), Tunisia (Kreiter et al. 2002),
Yemen (Ueckermann 1996). Asia: China (Wu 1980), Cyprus (Amitai 1992), Iran (Rahmani et al. 2010), Israel
(Swirski & Amitai 1965), Japan (Ehara 1972), Jordan (Allawi 1991), Oman (Hountondji et al. 2010), Saudi Arabia
(Al-Atawi 2011b), South Korea (Ryu 1997), Syria (Barbar 2013), Thailand (Oliveira et al. 2012), Turkey (Swirski
& Amitai 1982). Europe: England (Hughes 1948), Finland (Tuovinen 1993), France (Kreiter et al. 2000), Georgia
(Wainstein & Vartapetov 1973), Germany (Karg 1965), Greece (Papaioannou-Souliotis 1981), Italy (Athias-Henriot
1961), Latvia (Petrova et al. 2000), the Netherlands (van de Vrie 1963), Norway (Denmark & Edland 2002), Portu-
gal (Ferreira & Carmona 1994), Russia (Meshkov 1999), Spain (Ragusa & Athias-Henriot 1983), Sweden (Steeghs
et al. 1993), Ukraine (Wainstein & Shcherbak 1972). North America: USA (Denmark & Evans 2011). Oceania:
Australia (Waite & Gerson 1994), Hawaii (Denmark & Evans 2011). South America: Brazil (Moraes et al. 1993),
Chile (Ragusa & Vargas 2002).
Remarks. Beaulieu & Beard (2018) provided a detailed redescription and illustration of this species. They also
designated a neotype of the species. The species is distributed worldwide, including in Taiwan (Demite et al. 2020;
Liao et al. 2020). The species is considered a subtype III-e generalist predators that lives on soil/litter, and plants
(McMurtry et al. 2013). EPPO (2020) listed this species as a commercial biological control agent for thrips and
broad mite.
Material examined. Japan: one female (HAL095F295) from Vitis vinifera (Vitaceae), 3 Aug 2006; one female
(HAL095F308) from V. vinifera (Vitaceae), 8 Sept 2006.
Previous records. Africa: Egypt (Abo-Shnaf & Moraes 2014), Tunisia (Sahraoui et al. 2012). Asia: Armenia
(Arutunjan 1969), Azerbaijan (Abbasova 1972), China (Zhang et al. 2017), Iran (Asali Fayaz et al. 2011), Israel
(Swirski & Amitai 1985), Kazakhstan (Wainstein 1962), Saudi Arabia (Negm et al. 2012), Syria (Barbar 2014), Ta-
jikistan (Wainstein 1962), Turkey (Çobanoðlu 1991). Europe: France (Athias-Henriot 1966), Georgia (Wainstein &
Vartapetov 1973), Greece (Papadoulis & Emmanouel 1990), Hungary (Bozai 1980), Italy (Ragusa & Paoletti 1985),
Latvia (Salmane 1996), Moldova (Kolodochka 1980), Norway (Denmark & Edland 2002), Portugal (Espinha et al.
1998), Russia (Wainstein 1962), Serbia (Stojnić et al. 2002), Slovakia (Fend’a 2010), Spain (Iraola et al. 1997),
Switzerland (Airoldi et al. 1989), Ukraine (Livshitz & Kuznetsov 1972). North America: Mexico (Denmark & Ev-
ans 2011), USA (Congdon 2002). South America: Chile (Trincado et al. 2018).
Remarks. The species is distributed worldwide, except in Oceania. It is a native predator of spider mites and
thrips in Xinjiang Uygur Autonomous Region of China and is more adapted than nonnative species to hot and dry
climates (Zhang et al. 2017). It was recently showed to be promising predator of the whitefly Bemisia tabaci (Han
et al. 2020).
Material examined. Japan: three females one male (HAL095F372) from Vitis vinifera (Vitaceae), 26 Aug 2006;
one female (HAL095F307) from V. vinifera (Vitaceae), 31 Aug 2006; one female (HAL101B180) from Perilla
frutescens (Lamiaceae), 30 Mar 2012.
USA: four females (HAL099C147) from Hydrangea macrophylla (Hydrangeaceae), 2 Oct 2010; two females
(QAR101H033) from Fragaria ananassa (Rosaceae), 12 Sept 2011; two females (QAR101H035) from F. anan-
assa (Rosaceae), 19 Sept 2011; one female (QAR101H030) from Thymes sp. (Lamiaceae), 18 Oct 2011; one female
(QAR102H031) from Prunus persica (Rosaceae), 7 July 2013.
Material examined. Australia: one female (HAL098B609) from Hydrangea macrophylla (Hydrangeaceae), 28 Oct
2009.
Netherlands: one female (HAL101B179) from Helleborus sp. (Ranunculaceae), 30 Mar 2012; one female
(QAR101H008) from Trachelium caeruleum (Campanulaceae), 18 July 2011; one female (QAR101H011) from H.
macrophylla (Hydrangeaceae), 18 Aug 2011; one female (QAR102H021) from H. macrophylla (Hydrangeaceae),
16 June 2013; one female (QAR102H051) from H. macrophylla (Hydrangeaceae), 15 Sept 2013.
Previous records. Africa: Algeria (Athias-Henriot 1960), Egypt (Chant 1959), Morocco (McMurtry & Boun-
four 1989), Tunisia (Kreiter et al. 2004). Asia: Armenia (Arutunjan 1970), Azerbaijan (Gadzhiev & Abbasova
1965), Cyprus (Vassiliou et al. 2012), India (Sadana & Kanta 1971), Iran (Sepasgozarian 1977), Israel (Amitai &
Swirski 1978), Saudi Arabia (Al-Atawi 2011a), Turkey (Özman & Çobanoðlu 2001). Europe: Austria (Bohm 1960),
Belarus (Sidlyarevich 1966), England (Collyer 1956), Finland (Tuovinen 1993), France (Oudemans 1930b), Geor-
gia (Wainstein 1961), Germany (Dosse 1956), Greece (Papadoulis & Emmanouel 1991), Hungary (Bozai 1980),
Italy (Ragusa 1977), Latvia (Petrova et al. 2000), Moldova (Wainstein 1973), Netherlands (Chant 1959), Norway
(Denmark & Edland 2002), Poland (Wiackowski & Suski 1963), Portugal (Espinha et al. 1998), Russia (Meshkov
1999), Slovakia (Fend’a & Schniererová 2005), Slovenia (Bohinc & Trdan 2013), Spain (Escudero & Ferragut
1998), Sweden (Sellnick 1958), Switzerland (Chant 1959), Ukraine (Livshitz & Kuznetsov 1972). North America:
Canada (Nesbitt 1951), Mexico (Chant 1959), USA (Nesbitt 1951). Oceania: Australia (Beard 2001), New Zealand
(Chant 1959). South America: Chile (Ragusa & Vargas 2002).
Remarks. The species is distributed worldwide, but had no previous record in Taiwan. EPPO (2020) listed
this species as a commercially used biological control agent. This species has the lifestyle of Type III-e, generalist
predator, which has soil/litter habitats similar to N. barkeri. Both species considered effective natural enemies of
thrips and spider mites.
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 307
Neoseiulus imbricatus (Corpuz & Rimando, 1966)
Material examined. Thailand: one female (HAL100B196) from Asparagus officinalis (Asparagaceae), 21 Nov 2011.
Previous records. Asia: Azerbaijan (Abbasova 1972), China (Zhu & Chen 1980), India (Gupta 1986), Philip-
pines (Corpuz & Rimando 1966), Saudi Arabia (Alatawi et al. 2017), Thailand (Ehara & Bhandhufalck 1977).
Remarks. This species is distributed in Asia. No difference exists between our specimens and the paratype
specimen (Aca014 from UPLB-MNH). Wu et al. (2009) reported that the species is an effective natural enemy of
spider mites and tarsonemids in rice fields.
Material examined. USA: one female (HAL095F305) from Fragaria chiloensis (Rosaceae), 29 Aug 2006; one
female (HAL095F306) from F. chiloensis (Rosaceae), 29 Aug 2006; one female (HAL095F315) from F. chiloensis
(Rosaceae), 3 Oct 2006.
Previous records. North America: USA (Schuster & Pritchard 1963).
Remarks. The species is only distributed in the United States. Schuster & Pritchard (1963) reported the species
from Zostera marina on sea shore, a species habitat for phytoseiid mites. Denmark & Evans (2011) recorded addi-
tional habitat plants, including strawberry, and Malus sp. The collecting records of the species are all from Fragaria
chiloensis. The biological control potential of the species requires further confirmation.
Material examined. Japan: one female (HAL095F296) from Vitis vinifera (Vitaceae), 3 Aug 2006; one female
(HAL095F310) from Brassica rapa (Brassicaceae), 15 Sept 2006.
Thailand: one female (QAR102H022) from Lactuca sativa (Asteraceae), 20 June 2013.
Previous records. Africa: Cameroon (Zannou et al. 2006). Asia: China (Zhu & Chen 1980), Indonesia (Ehara
2002b), Japan (Ehara 1972), Saudi Arabia (Negm et al. 2012), South Korea (Ryu 1993), Taiwan (Tseng 1983),
United Arab Emirates (Negm 2014).
Remarks. The species is distributed in Africa and Asia. Ehara (1972) described this species from melons in
Japan. No difference was observed among these specimens and holotype (NSMT-AC13110 from NSMT). Liao et
al. (2020) reported a correlation between the species and tetranychid mites, but further experiments are required to
determine its biological control potential.
Material examined. Netherlands: three females one male (HAL095F302) from Symphoricarpos albus (Caprifo-
liaceae), 27 Aug 2006; three females one male (HAL095F303) from S. albus (Caprifoliaceae), 27 Aug 2006; one
female (HAL095F317) from Lactuca sativa var. capitata (Asteraceae), 12 Oct 2006.
Material examined. Japan: one female (HAL095F311) from Brassica rapa (Brassicaceae), 15 Sept 2006.
Previous records. Asia: China (Wu et al. 2009), Saudi Arabia (Negm et al. 2012), Taiwan (Tseng 1983), Unite
Arab Emirates (Negm 2014). Central America: Cuba (Ramos & Rodriguez 1999), Jamaica (Denmark & Muma
1978), Nicaragua (Rodríguez-Morell et al. 2013). North America: Mexico (Estebanes-Gonzale & Rodriguez-Na-
varro 1998), USA (Chant 1959). Oceania: Hawaii (Wainstein 1983). South America: Brazil (Denmark & Muma
1973).
Remarks. Liao et al. (2020) provided a detailed comparison between the species and Prop. mexicanus. Al-
though no significant morphological differences were found, further molecular study is needed in order to drive a
final conclusion.
The species is considered as lifestyle Type III-e, generalist predators from soil/litter habitats. Ho & Chen (2001)
proposed that this species have the potential to become natural enemies of thrips.
Material examined. Peru: one female (QAR102H029) from Asparagus officinalis (Asparagaceae), 21 July 2013.
Previous records. South America: Argentina (Guanilo et al. 2008a), Brazil (Ferla & Moraes 1998), Colombia
(Moraes & Mesa 1988), Ecuador (Moraes et al. 1991), Peru (Guanilo et al. 2008b).
Remarks. The species is distributed in South America. It is a generalist predator inhabiting natural vegetation
and crop fields (Moraes et al. 2004a). The biological control potential of the species requires further exploration.
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 309
Subfamily Typhlodrominae Scheuten
Material examined. USA: one female (HAL095F371) from Prunus persica (Rosaceae), 25 Sept 2006; six females
(HAL095F350) from M. pumila (Rosaceae), 9 Nov 2006; one female (HAL095F455) from M. pumila (Rosaceae),
15 Nov 2006; two females (HAL095F456) from M. pumila (Rosaceae), 15 Nov 2006; four females (HAL095F458)
from M. pumila (Rosaceae), 15 Nov 2006; two females (HAL095F459) from M. pumila (Rosaceae), 15 Nov 2006;
two females (TAL095F457) from Prunus persica (Rosaceae), 15 Nov 2006; four females (HAL095G068) from
M. pumila (Rosaceae), 30 Nov 2006; four females (HAL095G069) from M. pumila (Rosaceae), 30 Nov 2006;
three females (HAL095G070) from M. pumila (Rosaceae), 1 Dec 2006; seven females (HAL095G079) from M.
pumila (Rosaceae), 7 Dec 2006; four females (HAL095G081) from M. pumila (Rosaceae), 7 Dec 2006; seven
females (HAL095G799) from M. pumila (Rosaceae), 19 Dec 2006; nine females (HAL095G800) from M. pumila
(Rosaceae), 25 Dec 2006; three females (QAR101H022) from P. persica (Rosaceae), 19 Sept 2011; one female
(HAL101B056) from P. persica (Rosaceae), 21 Sept 2011; one female (QAR101H024) from P. persica (Rosaceae),
22 Sept 2011; one female (QAR101H025) from P. persica (Rosaceae), 22 Sept 2011; one female (QAR101H026)
from P. persica (Rosaceae), 22 Sept 2011; three females (HAL101B059) from P. persica (Rosaceae), 28 Sept 2011;
one female (QAR102H034) from P. persica (Rosaceae), 14 Aug 2013; one female (QAR102H046) from P. per-
sica (Rosaceae), 12 Sept 2013; one female (QAR102H047) from P. persica (Rosaceae), 12 Sept 2013; one female
(QAR102H053) from P. persica (Rosaceae), 26 Sept 2013.
Previous records. Africa: South Africa (Denmark 1982). Asia: China (Wu et al. 1997), Israel (Denmark 1982),
Jordan (Allawi 1991), South Korea (Ryu & Lee 1992), Taiwan (Chant & Yoshida-Shaul 1984). Europe: Austria (El-
Borolossy 1989), Greece (Ragusa Di Chiara et al. 1995), Netherlands (Hoying & Croft 1977), Russia (Denmark
1982). North America: Canada (Nesbitt 1951), Mexico (Denmark & Evans 2011), USA (Cunliffe & Baker 1953).
Oceania: Australia (Whitney & James 1996), New Zealand (Collyer 1964b). South America: Chile (Prado 1991),
Venezuela (Aponte & McMurtry 1993).
Remarks. The species is distributed worldwide (Demite et al. 2020). Denmark (1982) reported species distri-
bution in Taiwan without recording specimen information. Chant & Yoshida-Shaul (1984) reported the species to
be distributed in Taiwan based on one female specimen collected in 1950. Lo et al. (1986) reported that the species
was introduced to Taiwan but could not be kept in laboratory. All specimens were collected on peaches and apples
from the United States. The species has favorable efficacy for use in orchards. EPPO (2020) listed this species as a
commercially used biological control agent.
Material examined. South Africa: one female (HAL095G059) from Berzelia sp. (Bruniaceae), 27 Oct 2006; one
female (HAL095G060) from Berzelia sp. (Bruniaceae), 27 Oct 2006; one female (HAL095G051) from Brunia lae-
vis (Bruniaceae), 28 Oct 2006; one male (HAL095G059) from Berzelia sp. (Bruniaceae), 1 Dec 2006.
Previous records. Africa: South Africa (Van der Merwe 1968).
Remarks. The species has only been recorded in South Africa. All species were found from commodity of
South Africa in 2006, and it should be noted that the related habitat plants were only imported during this time.
Material examined. Thailand: two females one male (HAL100B063) from Rubus idaeus (Rosaceae), 30 June
2011.
Previous records. Europe: the Netherlands (Faraji 2006). North America: Canada (Nesbitt 1951), USA (Parrott
et al. 1906).
Remarks. The species has only been recorded in the Netherlands, Canada, and the United States. The speci-
mens were collected from commodity of Thailand, which had no previous record of the species. The distribution of
this species requires further confirmation.
Material examined. USA: one female (HAL095F312) from Prunus persica (Rosaceae), 16 Sept 2006; one female
(QAR101H020) from P. persica (Rosaceae), 16 Sept 2011; one female (QAR101H021) from P. persica (Rosaceae),
19 Sept 2011; one female (HAL101B058) from P. persica (Rosaceae), 23 Sept 2011; one female (QAR102H019)
from P. persica (Rosaceae), 8 June 2013; one female (QAR102H047) from P. persica (Rosaceae), 12 Sept 2013;
one female (QAR102H052) from P. persica (Rosaceae), 17 Sept 2013.
Japan: three females (QAR102H049) from Vitis vinifera (Vitaceae), 13 Sept 2013;
Previous records. Asia: Azerbaijan (Abbasova 1972), China (Wu 1988), Iran (Hajizadeh et al. 2002), Eu-
rope: Austria (El-Borolossy 1989), England (Collyer 1964), Latvia (Salmane & Petrova 2002), Lithuania (Pauriene
1970), Moldova (Beglyarov & Malov 1977), Norway (Evans & Edland 1998), Russia (Wainstein 1975), Slova-
kia (Jedlickova & Kolodochka 1994), Ukraine (Kolodochka 1978). North America: Canada (Downing & Moilliet
1971), USA (Schuster 1959). Oceania: Australia (Chant et al. 1978), New Zealand (Collyer 1964a).
Remarks. The species is distributed worldwide, except in Africa and South America. Most of the specimens
were found on peaches from the United States, except one was found on grapes from Japan.
Diagnosis. Female dorsal surface strongly reticulated, bearing 20 pairs of dorsal setae (including r3 and R1). All se-
tae smooth except Z5 is slightly serrated. Five pairs of solenostomes (gd2, gd4, gd6, gd8, gd9) visible on the dorsal
shield. Peritreme extending to level of seta j3. Sternal shield with two pairs of setae; ventrianal shield bearing four
pairs of pre-anal setae, with a pair of small rounded pre-anal solenostomes. Fixed digit of chelicera with four teeth;
movable digit with one tooth. Calyx of spermatheca cup-shaped with distal half lightly sclerotised. Leg IV with one
pair of macrosetae St IV; genu II with eight setae.
Female (n=4). A lightly sclerotised mite. Idiosomal setal pattern: 12A:8A/JV:ZV.
Dorsum (Fig. 1). Dorsal shield nearly oval, constricted at level of R1, strongly reticulated; 333 329 (314–339)
long (j1–J5 level) and 176 169 (159–176) wide at level of j6, 182 174 (163–182) wide at level of S4; five pairs
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 311
of solenostomes on dorsal shield, (gd2, gd4, gd6, gd8, gd9), thirteen pairs of lyrifissures, (id1, id2, id4, id6, idm2,
idm3, idm4, idm5, idm6, is1, idl2, idl3, idl4); length of setae: j1 21 21 (20–22), j3 24 27 (24–29), j4 15 17 (15–20),
j5 15 16 (15–17), j6 17 20 (17–23), J2 20 22 (20–25), J5 11 9 (6–11), z2 18 20 (18–22), z3 28 27 (26–28), z4 22
25 (22–26), z5 17 19 (17–22), Z4 34 33 (31–35), Z5 58 57 (54–59), s4 27 30 (27–33), s6 27 32 (27–35), S2 30 32
(30–34), S4 28 30 (28–31), S5 27 26 (25–27), r3 33 31 (28–33), R1 23 26 (23–29). All setae smooth, sharp pointed,
except for Z5 slightly serrated.
Peritreme (Figs 1, 2). Peritreme extending to level of seta j3; peritremal shield lightly sclerotised, with one pair
of solenostomes (gd3) and one pair of lyrifissures (id3).
FIGURES 1–4. Typhlodromus (Anthoseius) ueckermanni sp. nov., Female. 1. dorsal shield; 2. ventral idiosoma; 3. chelicera;
4. spermatheca.
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 313
Venter (Fig. 2). Sternal shield smooth, posterior margin irregular, much wider than long, 54 53 (49–58) long, 62
66 (62–71) wide, with two pairs of setae st1 27 25 (22–27), st2 27 24 (20–27), and two pairs of lyrifissures (pst1,
pst2). Sternal seta st3 23 23 (21–24) on separated platelets. Exopodal shield at coxae II–IV. Metasternal platelets
tear-shaped, with one pair of metasternal setae, st4 25 26 (25–28), and one pair of lyrifissures (pst3). Genital shield
smooth, truncate posteriorly, with one pair of genital setae st5 22 21 (18–22), 60 61 (57–66) wide at level of genital
setae. Distances between st1–st1 51 51 (48–52), st2–st2 51 55 (51–58), st5–st5 58 54 (50–58). Ventrianal shield
pentagonal with waist slightly below JV2 level, reticulated, 104 104 (96–115) long, 83 81 (79–83) wide at level of
ZV2, 73 74 (71–77) wide at level of anus; with four pairs of pre-anal setae, JV1 17 17 (17–19), JV2 15 13 (10–15),
JV3 12 15 (12–20), ZV2 13 16 (13–19), solenostomes gv3 small and rounded; Pa 13 12 (11–13), Pst 11 12 (11–15)
on shield. Setae JV4 18 16 (13–19), JV5 53 49 (47–53), ZV1 17 18 (14–21), ZV3 9 12 (9–14) on interscutal mem-
brane. All setae smooth, sharp pointed. Two metapodal plates: primary plates 25 27 (25–29) long, secondary plates
4 4 (3–4) wide, 11 10 (11–13) long, 1 2 (1–2) wide.
Spermatheca (Fig. 4). Calyx bell-shaped and elongated, flaring distally, with apical half thick (sclerotised), 17
18 (17–20) long, 9 11 (9–13) wide, atrium incorporated within the calyx, major duct broad.
Chelicera (Fig. 3). Movable digit 26 27 (26–27) long, with one tooth; fixed digit 25 26 (25–27) long, with four
teeth and pilus dentilis.
Legs (Figs 5–8). Complement of setae on coxae I IV: 2 2 2 1. Chaetotaxy (femur to basitarsus): leg I, 2 3/1 2/2
2, 2 2/1 1/1 2, 2 2/1 2/1 2, 1 1/0 1/0 1; leg II, 2 3/1 2/1 1, 2 2/1 2/0 1, 1 1/1 2/1 1, 1 1/0 1/0 1; leg III, 1 2/1 1/0 1, 1
2/1 2/0 1, 1 1/1 2/1 1, 1 1/0 1/0 1; leg IV, 1 2/1 1/0 1, 1 2/1 2/0 1, 1 1/1 2/0 1, 1 1/0 1/0 1. Macrosetae: St IV (pd) 33
30 (28–33). Macrosetae setiform.
Type specimens. Female Holotype: South Africa: one female (HAL095G078) from Rosmarinus officinalis
(Lamiaceae), 7 Dec 2006 (NTU). Paratypes: three females (HAL095G078) data same with holotype (NTU).
Etymology. The specific name “ueckermanni“ refers to Prof. Eddie Ueckermann, who provided detailed com-
parison of this species and T. (A.) lootsi.
Previous records. Asia: South Africa (present study).
Remarks. The new species was compared with all known species of the subgenus Anthoseius based on Phy-
toseiidae Database (Demite et al. 2020) and electronic polytomous key (Hernandes et al. 2012). Among them, it
shows a close affinity to T. (A.) aktherecus (Kolodochka), T. (A.) bagdasarjani Wainstein & Arutunjan, T. (A.)
georgicus Wainstein, T. (A.) involutus Livshitz & Kuznetsov, T. (A.) karaisaliensis Döker & Kazak, T. (A.) kerkirae
Swriski & Ragusa, T. (A.) lootsi Schultz, T. (A.) ponticus (Kolodochka), T. (A.) recki Wainstein, T. (A.) rhenanus
(Oudemans), T. (A.) salvia (Kolodochka), T. (A.) spiralis (Wainstein & Kolodochka), and mostly based on the shape
of calyx of spermatheca and general apperance. Differences between T. (A.) ueckermanni sp. nov. and related spe-
cies are given in Table 1.
Material examined. Australia: one female (HAL095G077) from Banksia sp. (Proteaceae), 7 Dec 2006.
Previous records. Oceania: Australia (Schicha 1978)
Remarks. The species has only been recorded in Australia. Schicha (1978) reported that the species fed on
an eriophyid mite colony from Ficus carica. He also reported that this species was collected from various parts of
Australia.
Material examined. France: one female (QAR102H041) from Thymes sp. (Lamiaceae), 6 Sept 2013.
Previous records. Africa: Algeria (Athias-Henriot 1960), Tunisia (Kreiter et al. 2002). Asia: Armenia (Arutun-
Material examined. USA: one female (HAL095G065) from Thymus vulgaris (Lamiaceae), 2 Nov 2006.
Netherlands: one female (HAL098B507) from Rosmarinus officinalis (Lamiaceae), 2 Oct 2009; one female
(HAL100B067) from R. officinalis (Lamiaceae), 19 May 2011; six females (HAL101B182) from R. officinalis
(Lamiaceae), 30 Mar 2012.
Previous records. Africa: Algeria (Athias-Henriot 1957b), Tunisia (Kreiter et al. 2002). Asia: Azerbaijan (Gad-
zhiev & Abbasova 1965), Cyprus (Georghiou 1959), India (Narayanan & Ghai 1961), Iran (Khalil-Manesh 1973),
Israel (Swirski & Amitai 1961), Kazakhstan (Andreeva 1966), Syria (Barbar 2013), Turkey (Çobanoðlu 1989a). Eu-
rope: Belarus (Sidlyarevich 1966), Belgium (Nesbitt 1951), Denmark (Hansen & Johnsen 1986), England (Nesbitt
1951), Finland (Tuovinen & Rokx 1991), France (Nesbitt 1951), Germany (Oudemans 1905a), Greece (Papadoulis
et al. 2009), Hungary (Kropczynska & Jenser 1968), Italy (Gunthart 1960), Latvia (Salmane & Petrova 2002),
Moldova (Kolodochka 1980b), Montengro (Mijuskovic & Tomasevic 1975), Netherlands (Miedema 1987), Nor-
way (Edland 1987), Poland (Suski 1961), Portugal (Carmona 1962), Russia (Beglyarov 1957), Serbia (Radivojević
& Petanović 1984), Slovakia (Fend’a 2010), Slovenia (Bohinc & Trdan 2013), Spain (Iraola et al. 1997), Sweden
(Steeghs et al. 1993), Switzerland (Gunthart 1960), Ukraine (Kolodochka 1974). North America: Canada (Nesbitt
1951), USA (Nesbitt 1951). South America: Brazil (Diehl et al. 2012);
Remarks. The species is distributed worldwide, except in Oceania. Chant (1959) reported the species after
Oudemans (1905) described it. Subsequent researchers provided conflicting opinions on the species status. Evans
& Momen (1988) examined the type materials of the species and related species and provided a detailed redescrip-
tion.
Material examined. Japan: one female (HAL100B195) from Vitis vinifera (Vitaceae), 10 Nov 2011.
Previous records. Asia: China [Hong Kong (Swirski & Shechter 1961)], Iran (Khalil-Manesh 1973), Japan
(Ehara 1959), South Korea (Ryu & Ehara 1990). Europe: Russia (Sapozhnikova 1966).
Remarks. The species is distributed in Asia and Europe. Kishimoto et al. (2013) reported that the species is
considered as lifestyle Type III, generalist predator that could become an effective predator of spider mites and eri-
ophyoid mites in field.
PHYTOSEIIDAE FROM TAIWAN Zootaxa 4927 (3) © 2021 Magnolia Press · 315
TABLE 1. Differences between Typhlodromus (Anthoseius) ueckermanni sp. nov. and related species.
Peritreme length solenostomes on dorsal shield Reticulation of dorsal shield gv3 No. of teeth on FD/MD* Morphology of St IV
a
ueckermanni j3 level 5 reticulated present 4/1 setiform
b
aktherecus j3 level 5 smooth present 3/1 setiform
c
bagdasarjani z3 level 5 reticulated absent 2/0 shovel-shaped with expanded blade
d
involutus j1level 5 reticulated present 4/2 setiform
karaisaliensis e j3 level 3 reticulated present 3/1 setiform
kerkirae f j3 level 5 reticulated present 3/1 setiform
g
lootsi j1level 3 reticulated absent 3/1 setiform
h
ponticus j1level 4 smooth present 3/1 shovel-shaped with expanded blade
recki i j3-z2 level 3 reticulated present 4/1 setiform
j
rhenanus j1level 5 reticulated present 3/1 setiform
k
salvie j1level 5 smooth present 4/1 setiform
TABLE 1. (Continued)
j3 Z4 Z5 S2 S4 S5 St IV
a
ueckermanni 24 27 (24–29) 34 33 (31–35) 58 57 (54–59) 30 32 (30–34) 28 30 (28–31) 27 26 (25–27) 33 30 (28–33)
b
aktherecus 34 41 54 41 41 54 30
c
bagdasarjani 31–35 53–55 68–73 41–44 35–38 29–32 49–54
involutus d 27–30 38 50 ? 27–30 15 55
e
karaisaliensis 48 (38–43) 52 (50–53) 69 (68–70) 51 (50–53) 43 (40–45) 38 (35–40) 34 (33–35)
f
kerkirae 28 (27-30) 33 (32-35) 51 (48-53) 32 (30-33) 32 (30-33) 28 (27-30) kerkirae
g
lootsi 27 61 53 47 47 48 short
h
ponticus 27 32 54 29 27 ? 27
recki i 28 (26–29) 31 (28–33) 50 (46–52) 31 (29–34) 32 (31–34) 25 (23–26) 31 (28–32)
rhenanus j 22–24 32–36 49–55 ? 33–36 29–33 18–19
k
salvie 22 31 50 31 ? ? 27
l
spiralis 28 42 58 33 31 ? 36
a
present study; b Kolodochka (1979); c Asali Fayaz et al. (2017); d Livshitz & Kuznetsov (1972); e Döker et al. (2017); f Swirski & Ragusa (1976) and Papadoulis et al. (2009); g
Schultz (1972); h Kolodochka (1992); i Rahmani et al. (2010); j Evans & Momen (1988); k Kolodochka (1979); lWainstein & Kolodochka (1974). * Teeth number of fixed digit and
movable digit of chelicerae.
LIAO ET AL.
Typhlodromus (Typhlodromus) pyri Scheuten, 1857
Material examined. Netherlands: one female (HAL100B198) from Rosmarinus officinalis (Lamiaceae), 3 Nov
2011.
Previous records. Africa: Egypt (El-Badry 1967). Asia: Azerbaijan (Abbasova 1970), Saudi Arabia (Fouly &
Al-Rehiayani 2011), Turkey (Çobanoðlu 1991). Europe: Austria (El-Borolossy 1989), Belarus (Sidlyarevich 1966),
Belgium (Chant et al. 1974), Croatia (Tixier et al. 2010), Czech Republic (Hluchy et al. 1991), Denmark (Chant
et al. 1974), England (Chant 1959), Finland (Tuovinen 1993), France (Rambier 1974), Germany (Scheuten 1857),
Greece (Swirski & Ragusa 1977), Hungary (Kropczynska & Jenser 1968), Italy (Castagnoli & Liguori 1986), Mol-
dova (Wainstein 1973), Montenegro (Mijuskovic & Tomasevic 1975), Netherlands (Van de Vrie 1963), Norway
(Edland 1987), Poland (Wiackowski & Suski 1963), Portugal (Carmona 1962), Russia (Meshkov 1999), Serbia
(Kropczynska & Petanović 1987), Slovakia (Jedlickova & Kolodochka 1994), Slovenia (Bohinc & Trdan 2013),
Spain (Pérez Otero & Mansilla Vázquez 1997), Sweden (Chant et al. 1974), Switzerland (Ragusa & Swirski 1976),
Ukraine (Kolodochka 1974). North America: Canada (Putman & Herne 1966), USA (Chant 1959). Oceania: Austra-
lia (Schicha 1987), New Zealand (Collyer 1964a). South America: Chile (Ragusa & Vargas 2002).
Remarks. The species is distributed worldwide, but only one specimens from the Netherlands has been found.
EPPO (2020) listed this species as a commercially used biological control agent. The species is considered as life-
style Type III-a, generalist predator that lives on pubescent leaves, and also has the ability to feed on fungi (McMur-
try et al. 2013).
Acknowledgements
We thank to BAPHIQ and inspectors who found the samples in plant quarantine. We also thank to E. Ueckermann
(Pretoria, South Africa), İ. Döker (CU, Turkey), S. F. Lin (NCHU, Taiwan), Y. Hsiao (CSIRO & ANU, Australia),
C. H. Chen (BAPHIQ, Taiwan), F. Y. Ning (BAPHIQ & NTU, Taiwan) for suggestions. Thank to H. Ono (NMNST,
Japan), L.A. Corpuz-Raros and J. Naredo (UPLB-MNH) for borrowing type specimens for comparison. Thanks to
S.F. Shiao (NTU, Taiwan) and all friends who provided great help after C.C. Ko passed away. Thanks to Wallace
Academic Editing for English editing of the draft. The study is supported by grants MOST 105-2621-B-002-002-
MY3 and MOST 108-2621-B-002-005-MY3 from the Ministry of Science and Technology, Taiwan.
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